Corallina officinalis on exposed to moderately exposed lower eulittoral rock

12-01-2005
Researched byDr Harvey Tyler-Walters Refereed byThis information is not refereed.
EUNIS CodeA1.122 EUNIS NameCorallina officinalis on exposed to moderately exposed lower eulittoral rock

Summary

UK and Ireland classification

EUNIS 2008A1.122Corallina officinalis on exposed to moderately exposed lower eulittoral rock
EUNIS 2006A1.122Corallina officinalis on exposed to moderately exposed lower eulittoral rock
JNCC 2004LR.HLR.FR.CoffCorallina officinalis on exposed to moderately exposed lower eulittoral rock
1997 BiotopeLR.ELR.FR.CoffCorallina officinalis on very exposed lower eulittoral rock

Description

Very exposed lower eulittoral rock on some shores supports a band of dense Corallina officinalis with low abundances of other turf-forming red algae such as Lomentaria spp., Mastocarpus stellatus, Ceramium spp. and Osmundea pinnatifida (=Laurencia pinnatifida), the red encrusting alga Callithamnion spp. and the brown alga Scytosiphon lomentaria. The coralline turf also creates a micro-habitat for small animals such as spirorbid worms. The brown alga Bifurcaria bifurcata and the barnacle Balanus perforatus may occur in the extreme south-west. This community usually forms a distinct band just above the kelp zone (EIR.Ala, EIR.Ala.Ldig or MIR.Ldig). (Information taken from the Marine Biotope Classification for Britain and Ireland, Version 97.06: Connor et al., 1997a, b).

Recorded distribution in Britain and Ireland

Recorded in various locations around Britain, although no records were found for Ireland.

Depth range

Lower shore, Mid shore

Additional information

None entered

Listed By

Further information sources

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JNCC

Habitat review

Ecology

Ecological and functional relationships

Coralline turf communities are described in detail by Hagerman (1968), Dommasnes (1968, 1969), Hicks (1985), Grahame & Hanna (1989), Crisp & Mwaiseje (1989), Bamber (1988) and Bamber & Irving ( 1993). The following information is based the above references and lists of species in the MNCR database (JNCC 1999).
  • Macroalgae including Corallina officinalis, Mastocarpus stellatus, Osmundea pinnatifida and Lomentaria articulata, provide primary productivity either directly to grazing fish and invertebrates or indirectly, to detritivores and decomposers, in the form of detritus and drift algae or as dissolved organic material and other exudates.
  • Macroalgal species compete for light, space and, to a lesser extent, nutrients, depending on the growth rates, size and reproductive pattern of each species. However, Corallina officinalis probably has a competitive advantage in wave exposed habitats due to their robust coralline fronds and resistant vegetative crustose bases (see Littler & Kauker, 1986).
  • Corallina officinalis provides substratum for spirorbid worms (e.g. Spirorbis corallinae), epiphytes and periphyton, depending on location, including microflora (e.g. bacteria, blue green algae, diatoms and juvenile larger algae), and interstices and refuges from predation for a variety of small invertebrates (see habitat complexity below).
  • Amphipods (e.g. Parajassa pelagica and Stenothoe monoculoides), isopods (e.g. Idotea pelagica and Jaera albifrons) and other mesoherbivores graze the epiphytic flora and senescent macroalgal tissue, which may benefit the macroalgal host, and may facilitate dispersal of the propagules of some macroalgal species (Brawley, 1992b; Williams & Seed, 1992). Mesoherbivores also graze the macroalgae but do not normally adversely affect the canopy (Brawley, 1992b).
  • Grazers of periphyton (bacteria, blue-green algae and diatoms) or epiphytic algae include harpacticoid copepods, small gastropods (e.g. Rissoa spp. and Littorina neglecta.
  • Macroalgal grazers include limpets e.g. Patella vulgata and Patella ulyssiponensis, juvenile blue-rayed limpets Helcion pelucidum, and gastropods such as Littorina saxatilis and Littorina neglecta.
  • Coralline algae are probably relatively grazing resistant (Littler & Kauker, 1984) and few species graze the corallines directly except perhaps chitons and limpets of the genus Tectura. Grazers probably benefit the coralline turf by removing epiphytic and ephemeral algae (e.g. Ulva), which could potentially smother the turf.
  • Suspension feeders include Semibalanus balanoides, the spirorbid Spirorbis corallinae, the sponge Halichondria panicea, juvenile bivalves and interstitial bivalves such as Lasaea adansoni and Turtonia minuta, and the tubiculous amphipod Parajassa pelagica.
  • Turbellarians, nematodes and halacarid mites are probably interstitial predators on other nematodes, mites, and harpacticoid copepods (Hicks, 1985).
  • When the biotope is covered by the tide, intertidal fish such as gobies, blennies and clingfish, and the juveniles of larger inshore fish are probably active predators of amphipods, isopod, ostracods and harpacticoid copepods. The physical complexity of the Corallina officinalis turf was reported to offer a refuge from predation for epiphytic invertebrates (Coull & Wells, 1983; Hicks, 1985). Choat & Kingett (1982) did not detect any significant effect on fish predation in exclusion experiments. In harpacticoid copepods, although large numbers were consumed by fish little effect on the population resulted (Hicks, 1980). However, Hicks (1985) noted that considerable evidence of predators regulating prey abundance was available.
  • The brittlestar Amphipholis squamata probably is a detritivore within the turf.

Seasonal and longer term change

Red algal turf declines in abundance during the winter months, partly due to die back and abrasion during winter storms. For example, Seapy & Littler (1982) noted that the cover of Corallina officinalis var. chilensis declined in the winter months, growing back in summer and developing a dense cover in autumn in California. Littler et al. (1979) reported a autumn maximum in cover of Corallina officinalis var. chilensis and a summer minimum in cover in San Clement Island, California. In Denmark, fronds of Corallina officinalis were reported to cease growing in summer, sloughed in autumn, and new fronds initiated from crustose, perenniating bases in late winter (Rosenvinge, 1917; cited in Johanssen, 1974). However, in the Bristol Channel, Bamber & Irving (1993) noted that the biomass of Corallina officinalis increased steadily through spring and summer and began to decline after July. Mastocarpus stellatus (as Gigartina stellata) was reported have a perennial holdfast, losing many erect fronds in winter, which grow back in spring (Dixon & Irvine, 1977). Osmundea pinnatifida also shows seasonal variation in growth, expanding its perennial holdfast in June to September, and producing erect fronds from October onwards reaching a maximum in February to May (Maggs & Hommersand, 1993).

Choat & Kingett (1982) reported that the abundance of amphipods in a New Zealand coralline turf habitat peaked in summer and declined to a low in winter, while polychaetes showed a peak of abundance in winter decreasing in summer. But ostracods showed a relatively low abundance throughout the sampling period (Choat & Kingett, 1982). Bamber (1993) examined coralline turf dominated runoffs in the Bristol Channel, and noted that the amphipod Melita palmata and the brittlestar Amphipholis squamata recruited after the summer growth of the coralline turf reaching a peak abundance in autumn. But the small isopod Jaera albifrons recruited to the turf in late winter and the polychaete Platynereis dumerilii showed an erratic pattern of abundance (Bamber & Irving, 1993). However, Bamber & Irving (1983) noted considerable variation in seasonal abundance between sites (runoffs) on the same shore.

Habitat structure and complexity

This biotope occurs in very wave exposed conditions on horizontal, steep or vertical bedrock subject to wave crash and is composed of species tolerant of wave action. The biotope may develop below the lower limit of the barnacle or mussel belts in wave exposed conditions.
  • Corallina officinalis forms a dense carpet or turf on the bedrock and with increasing wave exposure may grow as a cushion like or compact turf (Dommasnes, 1968; Johansen, 1974; Irvine & Chamberlain, 1994).
  • Other red algae occur in low abundance depending on wave exposure with Mastocarpus stellatus being the most tolerant, Osmundea pinnatifida slightly less tolerant, while Lomentaria articulata and Palmaria palmata favour shaded or overhanging surfaces. Shaded overhangs may also support Plumaria elegans, Ptilota plumosa and Cladophora rupestris (Lewis, 1964).
  • Depressions filled with Osmundea pinnatifida and Corallina officinalis may also support the olive-brown bulbous seaweed Leathesia difformis (Lewis, 1964).
  • Large macroalgae such as Himanthalia elongata typically occur at low abundance, their long thongs lying over the coralline turf.
  • The interstices formed by the branches of Corallina officinalis support a diverse epiphytic fauna (Dommasnes, 1968, 1969; Hagerman, 1968; Hicks & Coull, 1983; Hicks, 1985; Bamber, 1988; Crisp & Mwaiseje, 1989; Grahame & Hanna, 1989; Bamber & Irving, 1993). The species diversity and abundance of the epiphytic fauna depends the percentage cover of turf, wave exposure, the size of the interstices within the turf, and the build up of sediment. In wave exposure, the build up of sediment is likely to be limited and the close compact, cushion growth form may reduce the diversity of the infauna but provide a better refuge from predation for harpacticoid copepods and ostracods (Dommasnes, 1968, 1969; Seapy & Littler, 1982; Choat & Kingett, 1982; Hicks & Coull, 1983; Hicks, 1985).
  • In wave exposed conditions, tubiculous amphipods and isopods are represented by species with well developed claws or gnathopods and strong stout legs and bodies, e.g. the isopods Idotea pelagica and Jaera albifrons, and the amphipods Parajassa pelagica, although Stenothoe monoculoides, Apherusa jurinei and the isopod Ianiropsis breviremis occur irrespective of wave exposure (Dommasnes, 1986, 1969).
  • Corallina officinalis provides a substratum for small spirorbids e.g. Spirobis corallinae, which is only found on Corallina officinalis. Increasing density of Spirorbis corallinae was shown to increase the species richness of the epiphytic fauna, especially small species such as Stenothoe monocloides (Crisp & Mwaiseje, 1989) but with increasing wave exposure, the spirorbid is found within the Corallina officinalis turf rather than at its tips and was reported to be absent from the 'most wave exposed' sites (Grahame & Hanna, 1989).
  • Wave exposed coralline turf also reported to support Foraminifera, Turbellaria, nematodes, polychaetes (e.g. Platynereis dumerilii and Perinereis cultrifera), the tanaid Tanais cavolinii, halacarid mites, gastropods (e.g. Littorina neglecta, Littorina saxatilis, and Rissoa spp.), juvenile bivalves (e.g. Mytilus edulis, Musculus discors), interstitial bivalves (e.g. Lasaea adansoni and Turtonia minuta) and the small brittlestar Amphipholis squamata (Hagerman, 1968; Dommasnes, 1968, 1969; Bamber & Irving, 1993).
  • In gaps in the turf, the surface of the bedrock may be covered with encrusting coralline algae and barnacles such as Semibalanus balanoides, and patrolled by limpets (e.g. Patella ulyssiponensis).
  • Productivity

    Little information concerning the productivity of coralline turf communities was found. The red algae, algal epiphytes and periphyton provide primary productivity to grazers, while their spores and phytoplankton provide primary productivity to suspension feeders. Bamber & Irving (1993) reported that Corallina officinalis reached a biomass of up to 3.3-6.7 kg/m². Littler et al. (1979) determined the total daily productivity of an intertidal algal population in California, which peaked in autumn at 1.22 gC fixed /m²/day, and declined in winter to a spring low of 0.47 gC fixed /m²/day. Blue-green algae, Corallina officinalis var. chilensis and Egregia menziesii contributed 76% of the total community primary productivity (Littler et al., 1979).

    Secondary productivity of the invertebrate fauna may be high and coralline turf may support high abundances of invertebrates. For example, Choat & Kingett (1982) recorded the following numbers of epiphytic fauna: amphipods 1038 / 0.01m²; ostracods 219 /0.01m², and polychaetes 134 /0.01m².

    Recruitment processes

    Corallina officinalis has isomorphic sexual (gametophyte) and asexual (sporophyte) stages (see MarLIN review). Settled tetraspores develop into a perennial crustose base, from which the upright, articulate fronds develop. Sporeling formed within 48hrs, a crustose base within 72hrs, fronds being initiated after 3 weeks and the first intergeniculum (segment) formed within 13 weeks (Jones & Moorjani, 1973). Settlement and development of fronds is optimal on rough surfaces but settlement can occur on smooth surfaces (Harlin & Lindbergh, 1977; Wiedeman, pers comm.). Corallina officinalis settled on artificial substrata within 1 week of their placement in the intertidal in New England summer suggesting that recruitment is high (Harlin & Lindbergh, 1977).

    The propagules of most macroalgae tend to settle near the parent plant (Schiel & Foster, 1986; Norton, 1992; Holt et al., 1997). For example, the propagules of fucales are large and sink readily and red algal spores and gametes and immotile. Norton (1992) noted that algal spore dispersal is probably determined by currents and turbulent deposition (zygotes or spores being thrown against the substratum). For example, spores of Ulva sp. (as Ulva) have been reported to travel 35km, Phycodrys rubens 5km and Sargassum muticum up to 1km, although most Sargassum muticum spores settle within 2m. The reach of the furthest propagule and useful dispersal range are not the same thing and recruitment usually occurs on a local scale, typically within 10m of the parent plant (Norton, 1992). In clearance studies in the subtidal Kain (1975) noted that on a single block cleared every two months, most biomass belonged to Rhodophyceae in winter, Phaeophyceae in spring and Chlorophyceae in late summer, and concluded that recruitment was dependant on spore availability. For example, spore production in Mastocarpus stellatus is maximum between September to December (Dixon & Irvine, 1977), spores of Osmundea pinnatifida are present in October and December to June (Maggs & Hommersand, 1993), while the spores of Lomentaria articulata are available all year round with a peak in summer (Irvine, 1983).

    Recruitment of Patella vulgata fluctuates from year to year and from place to place (Bowman, 1981). Fertilization is external and the larvae are pelagic for up to two weeks before settling on rock at a shell length of about 0.2mm. Winter breeding occurs only in southern England, in the north of Scotland it breeds in August and in north-east England in September. Reproduction is probably similar in Patella ulyssiponensis, except that it may be a protandrous hermaphrodite, spawning in October in south-west Ireland (Fish & Fish, 1996). The larvae of the blue-rayed limpet Helcion pellucidum settle on encrusting corallines and migrate to Mastocarpus stellatus as they grow and finally to Laminaria spp. via Himanthalia elongata (McGrath, 1992; see MarLIN review).

    Barnacle recruitment can be very variable because it is dependent on a suite of environmental and biological factors, such as wind direction and success depends on settlement being followed by a period of favourable weather. Long term surveys have produced clear evidence of barnacle populations responding to climatic changes. During warm periods Chthamalus spp. predominate, whilst Semibalanus balanoides does better during colder spells (Hawkins et al., 1994). Release of Semibalanus balanoides larvae takes place between February and April with peak settlement between April and June.

    Many species of mobile epifauna, such as polychaetes have long lived pelagic larvae and/or are highly motile as adults. Gammarid amphipods brood their embryos and offspring but are highly mobile as adults and probably capable of colonizing new habitats from the surrounding area (e.g. see Hyale prevosti review). Similarly, isopods such as Idotea species and Jaera species brood their young. Idotea species are mobile and active swimmers and probably capable to recruiting to new habitats from the surrounding area by adult migration. Jaera albifrons, however, is small and may take longer to move between habitats, and Carvalho (1989) suggested that under normal circumstances movement was probably limited to an area of less than 2m. Hicks (1985) noted that epiphytic harpacticoid copepods lack planktonic dispersive larval stages but are active swimmers, which is therefore the primary mechanism for dispersal and colonization of available habitats. Some species of harpacticoids are capable to moving between low and mid-water levels on the shore with the tide, while in other colonization rates decrease with increasing distance form resident population. Overall immigration and in situ reproduction were thought to maintain equilibrium populations exposed to local extinction, although there may be local spatial variation in abundance (see Hicks, 1985).

    The small littorinids Littorina saxatilis and Littorina neglecta are ovoviviparous, releasing miniature adults. Therefore, local recruitment is probably good, whereas long distance recruitment is probably poor. The interstitial bivalve Lasaea adansoni also broods its eggs, releasing miniature adults. However, Martel & Chia (1991b) reported bysso-pelagic or mucus rafting in small bivalves and gastropods in the intertidal, and suggested that drifting may be an effective mean of dispersal at the local scale, even for species that produce miniature adult offspring. The gastropod Rissoa parva lays eggs capsules, from which hatch veliger larvae with a prolonged pelagic life and potentially good dispersal capability (Fish & Fish, 1996).

    Time for community to reach maturity

    The epiphytic species diversity of the coralline turf is dependant on the Corallina officinalis cover and its growth form (Dommasnes, 1968, 1969; Seapy & Littler, 1982; Crisp & Mwaiseje, 1989). Corallina officinalis was shown to settle on artificial substrata within one week of their placement in the intertidal in New England summer suggesting that recruitment is high (Harlin & Lindbergh, 1977). New fronds of Corallina officinalis appeared on sterilised plots within six months and 10% cover was reached with 12 months (Littler & Kauker, 1984). In experimental plots, up to 15% cover of Corallina officinalis fronds returned within 3 months after removal of fronds and all other epiflora/fauna (Littler & Kauker, 1984). Bamber & Irving (1993) reported that new plants grew back in scraped transects within 12 months, although the resistant crustose bases were probably not removed. New crustose bases may recruit and develop quickly the formation of new fronds from these bases and recovery of original cover may take longer. Once a coralline turf has developed it will probably be colonized by epiphytic invertebrates such as harpacticoids, amphipods and isopods relatively quickly from the surrounding area. Therefore, the biotope would be recognizeable once the coralline turf has regrown, which is likely to be within a few months if the resistant crustose bases remain. Recruitment of red algae is probably equally rapid, and once the algal turf has developed most of the epiphytic invertebrates would colonize quickly, although some species e.g. small brooding gastropods would take longer.

    Additional information

    None entered

Preferences & Distribution

Recorded distribution in Britain and IrelandRecorded in various locations around Britain, although no records were found for Ireland.

Habitat preferences

Depth Range Lower shore, Mid shore
Water clarity preferences
Limiting Nutrients No information found
Salinity Full (30-40 psu)
Physiographic Open coast
Biological Zone Lower eulittoral
Substratum Bedrock
Tidal Moderately Strong 1 to 3 knots (0.5-1.5 m/sec.), Very Weak (negligible), Weak < 1 knot (<0.5 m/sec.)
Wave Exposed, Moderately exposed, Very exposed
Other preferences Wave exposed conditions

Additional Information

This biotope is characteristic of wave exposed headlands and the open coast on steep to vertical slopes exposed to the full impact of wave crash or horizontal scarps from which water drains slowly (Lewis, 1964; Connor et al., 1997b). The ELR.Coff community often forms a distinct band below mussel or barnacle dominated communities and above the kelp belt, the coralline turf often extending into the kelp belt, e.g. EIR.Ala (Lewis, 1964; Connor et al., 1997b).

Species composition

Species found especially in this biotope

Rare or scarce species associated with this biotope

-

Additional information

The MNCR recorded 104 species within this biotope, although not all species occurred in all records of the biotope (JNCC, 1999). Detailed lists of the fauna of coralline turfs are given by Hagerman (1968), Dommasnes (1968, 1969), Hicks (1985), Grahame & Hanna (1989), Crisp & Mwaiseje (1989), and Bamber (1988, 1993).

Sensitivity reviewHow is sensitivity assessed?

Explanation

Corallina officinalis is the dominant characterizing species within the biotope and provides substratum and refuges for a diverse fauna. Therefore, the community is dependant on the presence of Corallina officinalis and it has been included as key structuring. Epiphytic grazers, such as amphipods, isopods small gastropods probably keep the turf free of epiphytic algae and are important functional species. Reference has been made to reviews of Hyale prevosti to represent the sensitivity of amphipods and small crustaceans. Similarly reference was made to Ahnfeltia plicata and Chondrus crispus to represent the sensitivity of characterizing intertidal red algae.

Species indicative of sensitivity

Community ImportanceSpecies nameCommon Name
Key structuralCorallina officinalisCoral weed
Important functionalGammaridaeGammarid amphipods
Important functionalIdotea pelagicaAn isopod

Physical Pressures

 IntoleranceRecoverabilitySensitivitySpecies RichnessEvidence/Confidence
High High Moderate Major decline High
Removal of the substratum would result in loss of the coralline turf and its associated community. Therefore an intolerance of high has been recorded. Recoverability is likely to be high (see additional information below).
Intermediate Very high Low Minor decline High
Seapy & Littler (1982) examined the effects of smothering of the intertidal with a layer of sediment in Santa Cruz, California after unusually heavy rainfall. In the 3 months that followed the total macrophyte cover decreased from 45.3 to 37.3% while macroinvertebrates, especially barnacles, were adversely effected declining from 15.8 to 6.5% cover. The Corallina spp. turf suffered a substantial decline, while the taller red alga Gigartina canaliculata was relatively unaffected. But in the following 6 months, the die back of higher shore species, e.g.. barnacles and Pelvetia spp, allowed the coralline turf and associated red algae to expand up the shore, and Corallina officinalis var. chilensis became the primary cover organism (Seapy & Littler, 1982). In ELR.Coff smothering may adversely affect the resident barnacle species. Smothering sediment will probably fill the interstices in the coralline turf excluding mobile invertebrates and interfering with feeding and respiration is tubicolous amphipods and worms, so that species diversity is likely to decrease. But in the wave exposed conditions characterized by this biotope, smothering is likely to be short lived. Therefore, an intolerance of intermediate has been recorded, and recovery is likely to be very high (see additional information below).
Intermediate Very high Low Minor decline Low
Corallina spp. accumulate more sediment than any other alga (Hicks, 1985). Hence an increase in suspended sediment is likely to accumulate in the coralline turf. A significant increase may result in smothering (see above). An accumulation of sediment within the turf may attract more sediment dwelling interstitial invertebrates such as nematodes, harpacticoids and polychaetes although, in this wave exposed habitat, accumulation of sediment is likely to be minimal. Increased suspended sediment is likely to result in increased scour, especially if the sediment is sand, which may adversely affect the fleshy red algae, and interfere with settling spores and recruitment if the factor is coincident with their major reproductive period. However, coralline algae, especially the crustose forms are thought to be resistant of sediment scour (Littler & Kauker, 1984), and will probably not be adversely affected at the benchmark level. Therefore, an increase in suspended sediment may reduce the epiphytic species diversity in the immediacy, and adversely affect the cover of fleshy red algae and an intolerance of intermediate has been recorded. Recoverability is likely to be very high (see additional information below).
Tolerant Very high Not sensitive* Decline Low
This community is unlikely to be dependant on suspended sediment. Although accumulated sediment within coralline turf habitats has been shown to increase the species diversity of the epiphytic fauna (see habitat complexity), in very wave exposed habitat the accumulated sediment is likely to be minimal. A reduction in suspended sediment will probably reduce the risk of scour, and reduce food availability for the few suspension feeding species in the biotope (e.g. barnacles and spirorbids if present). Therefore not sensitive has been recorded.
High High Moderate Major decline Moderate
Finely branched fronds or cushion-like turfs may hold water, reducing desiccation stress. Corallina officinalis inhabits damp or wet gullies and rock pools and does not inhabit the upper shore, suggesting that it is intolerant of desiccation. Desiccation risk is reduced in wave exposed habitats, and with increasing wave exposure the coralline turf may extend further up the shore. However, its upper limit is probably determined by competition, since Seapy & Littler (1982) found that die back of higher shore species allowed coralline species and associated foliose red algae to extend up the shoe. On moderately wave exposed shores ELR.Coff is probably restricted to gentle sloping or horizontal substrata that drain slowly.

Fronds of Corallina officinalis are highly intolerant of desiccation and do not recover from a 15% water loss, which might occur within 40 -45 minutes during a spring tide in summer (Wiedemann, 1994). An abrupt increase in temperature of 10 °C caused by the hot, dry 'Santa Anna' winds (between January and February) in Santa Cruz, California resulted in die back of several species of algae exposed at low tide (Seapy & Littler, 1982). Although fronds of Corallina spp. dramatically declined, summer regrowth resulted in dense cover by the following October, suggesting that the crustose bases survived. The red alga Gigartina canaliculata decreased at its upper limit but increased lower on the shore (Seapy & Littler, 1982). Severe damage was noted in Corallina officinalis as a result of desiccation during unusually hot and sunny weather in summer 1983 (an increase of between 4.8 and 8.5 °C above normal) (Hawkins & Hartnoll, 1985). Similarly, red algae such as Mastocarpus stellatus and Osmundea pinnatifida were damaged or killed at their upper limits. Hawkins & Harkin (1985) found that Corallina officinalis and encrusting corallines often die when their protective canopy of other algal species is removed. Overall, the evidence suggests that this community is likely to be highly intolerant of increased desiccation, equivalent to being raised one level on the shore, and would probably be lost. Recovery is likely to be rapid (see additional information below).

Intermediate Very high Low Minor decline Low
Bleached corallines were observed 15 months after the 1964 Alaska earthquake which elevated areas in Prince William Sound by 10 m. Similarly, increased exposure caused by upward movement of 15 cm due to nuclear tests at Armchitka Island, Alaska adversely affected Corallina pilulifera (Johansen, 1974). An increase in emergence is likely to result in decreased wetting and hence increased risk of desiccation. Therefore, the upper limit of this biotope is likely to be depressed and an intolerance of intermediate has been recorded. Recoverability is probably very high (see additional information below).
Tolerant* Not sensitive No change Low
A decrease in emergence will reduce the risk of desiccation, and increase the average wetness of the shore, potentially allowing the community to expand further up the shore. Therefore, not sensitive* has been recorded. However, it is also likely that the lower extent of the biotope would change to a kelp dominated community.
Tolerant Not relevant Not relevant Not relevant Low
This biotope occurs in moderately strong to very weak tidal streams. Water movement is probably an important structuring feature of the biotope as dense coralline turfs only develop on open bedrock in wave exposed conditions. Wave action is of greater importance than water flow in this biotope and it is unlikely that increased water flow will have an impact. Therefore, not sensitive has been recorded.
High Moderate Intermediate Rise Moderate
In low water flow, wave action is probably the most important cause of water movement within the biotope. Therefore, not relevant has been recorded.
Intermediate Very high Low Decline Moderate
Lüning (1990) reported that Corallina officinalis from Helgoland survived one week exposure to temperatures between 0 °C and 28 °C. An abrupt increase in temperature of 10 °C caused by the hot, dry 'Santa Anna' winds (between January -and February) in Santa Cruz, California resulted in die back of several species of algae exposed at low tide (Seapy & Littler, 1984). Although fronds of Corallina spp. dramatically declined, summer regrowth resulted in dense cover by the following October, suggesting that the crustose bases survived. Severe damage was noted in Corallina officinalis as a result of desiccation during unusually hot and sunny weather in summer 1983 (an increase of between 4.8 and 8.5 °C) (Hawkins & Hartnoll, 1985). Littler & Kauker (1984) suggested that the crustose base was more resistant of desiccation or heating than fronds.

Most of the other species within the biotope are distributed to the north and south of Britain and Ireland and unlikely to be adversely affected by long-term temperature change. But Hawkins & Hartnoll (1985) suggested that typical understorey red algae were susceptible to hot dry weather and that occasional damaged specimens of Palmaria palmata, Osmundea pinnatifida and Mastocarpus stellatus were observed after the hot summer of 1983.

It is likely that Corallina officinalis fronds are intolerant of abrupt short term temperature increase although they may not be affected by long term chronic change and the crustose bases are probably more tolerant than fronds. Epiphytic species will decline due to loss of coralline turf cover. Similarly, acute increases in temperature will probably reduce the cover of the characterizing red algae. Therefore, an intolerance of intermediate has been recorded, although recoverability in likely to be very high.

Low Immediate Not relevant Minor decline Low
Lüning (1990) reported that Corallina officinalis from Helgoland survived one week exposure to temperatures between 0 °C and 28 °C. New Zealand specimens were found to tolerate -4 °C (Frazier et al., 1988, cited in Lüning, 1990). Lüning (1990) suggested that most littoral algal species were tolerant of cold and freezing. For example, the photosynthetic rate of Chondrus crispus recovered after 3hrs at -20 °C but not after 6hrs exposure (Dudgeon et al., 1990). The photosynthetic rate of Mastocarpus stellatus higher on the shore fully recovered from 24hrs at -20 °C. Lüning reported that optimal growth and temperature were between 10-20 °C for Lomentaria articulata, consistent with their peak spore production in summer. Little information was found on the effects on the epiphytic fauna but all species are adapted to the extreme fluctuations in temperature that occur in the intertidal and are probably tolerant of acute temperature change. Overall, the biotope is unlikely to be adversely affected by long term temperature change at the benchmark level. Short term acute change my result in loss of a few individuals of more intolerant species e.g. Semibalanus balanoides and Patella species (see MarLIN reviews) but otherwise not adversely affect the biotope. Therefore an intolerance of low has been recorded.
Tolerant Not relevant Not relevant No change Low
Red algae and coralline algae especially are known to be shade tolerant and are common components of the understorey on seaweed dominated shores. Therefore, a decrease in light intensity is unlikely to adversely affect the biotope, especially as it is emersed regularly with the tide. Hence, not sensitive has been recorded.
Tolerant Not sensitive* No change Low
An increase in light intensity is unlikely to adversely affect the biotope. The community is regularly uncovered by the receding tide and experiences full sunlight, which may bleach the tips of algal fronds. Therefore, not sensitive has been recorded.
Not relevant Not relevant Not relevant Minor decline High
Wave exposure is an important determining factor in this biotope, removing competition from fleshy red algae and fucoids and allowing a dense coralline turf to develop. The biotope occurs in extremely wave expose to moderately wave exposed habitats so that any further increase in wave exposure is unlikely. Hence, not sensitive has been recorded. In records of ELR.Coff in moderately wave exposed habitats an increase in wave exposure may decrease the species richness of the epiphytic fauna.
High Moderate Intermediate Rise Moderate
Wave exposure is an important determining factor in this biotope, removing competition from fleshy red algae and fucoids and allowing a dense coralline turf to develop. The biotope occurs in extremely to moderately wave exposed habitats. A decrease in wave exposure from e.g. very exposed to moderately exposed may increase the abundance or fleshy red algae, and diversity of the epiphytic fauna and probably reduce the height and width of the coralline turf zone in the intertidal. But a decrease in wave exposure from e.g. exposed to sheltered would result in major changes in the community, probably favouring fucoid dominated biotopes and resulting in loss of the biotope as described. Therefore, an intolerance of high has been recorded. Recoverability could be high (see additional information below) but may only occur once the dominant species in the replacement biotope have died or been removed by wave action, which may take more than five years. Therefore, a recoverability of moderate has been recorded in this instance.
Tolerant Not relevant Not relevant No change Not relevant
None of the species in this biotope are know to respond to noise or vibration at the benchmark level, and live in a habitat that experiences the severe noise and turbulence caused by wave action.
Tolerant Not relevant Not relevant No change High
The mobile invertebrates are probably capable of responding to localized shading, experienced by the approach of a predator. But their visual acuity is likely to be low and they are unlikely to respond to visual disturbance at the benchmark level.
Intermediate Very high Low Decline Moderate
Abrasion by an anchor or mooring may remove some fronds of the foliose red algae and coralline turf, although most species would grow back from their remaining holdfasts. Trampling may be more damaging. For example, moderate (50 steps per 0.09 sq. metre) or more trampling on intertidal articulated coralline algal turf in New Zealand reduced turf height by up to 50%, and the weight of sand trapped within the turf to about one third of controls. This resulted in declines in densities of the meiofaunal community within two days of trampling. Although the community returned to normal levels within 3 months of trampling events, it was suggested that the turf would take longer to recover its previous cover (Brown & Taylor, 1999). Similarly, Schiel & Taylor (1999) noted that trampling had a direct detrimental effect on coralline turf species on the New Zealand rocky shore. At one site, coralline bases were seen to peel from the rocks (Schiel & Taylor 1999), although this was probably due to increased desiccation caused by loss of the algal canopy. The crustose base has nearly twice the mechanical resistance (measured by penetration) of fronds (Littler & Kauker, 1984). Brosnan & Cumrie (1994) also reported that foliose algae, e.g. Mastocarpus papillatus showed significant declines in cover in response to trampling, although recovery was rapid, probably from remaining holdfasts. Therefore, physical abrasion due to trampling is likely to result in a significant decline in the cover of the coralline turf, red algae and epiphytic fauna and an intolerance of intermediate has been recorded. The dominant algae are likely to recover rapidly by regrowth from remaining fronds and holdfasts, and epiphytic fauna will colonize the turf relatively quickly.
High High Moderate Major decline Moderate
The majority of the epiphytic fauna, such as the isopods, amphipods and harpacticoid copepods are highly mobile are unlikely to be adversely affected by displacement. But the dominant macroalgae are permanently attached to the substratum and if removed will be lost, resulting in loss of the biotope overall. If their holdfasts and bases are also removed then recovery will be prolonged but still relatively rapid.

Chemical Pressures

 IntoleranceRecoverabilitySensitivityRichnessEvidence/Confidence
High High Moderate Major decline Low
Smith (1968) reported that oil and detergent dispersants from the Torrey Canyon spill affected high water specimens of Corallina officinalis more than low shore specimens and some specimens were protected in deep pools. In areas of heavy spraying, however, Corallina officinalis was killed, and was affected down to 6m depth at one site, presumably due to wave action and mixing (Smith, 1968). However, regrowth of fronds had begun within 2 months after spraying ceased (Smith, 1968). O'Brien & Dixon (1976) suggested that red algae were the most sensitive group of algae to oil or dispersant contamination, possibly due to the susceptibility of phycoerythrins to destruction. They also report that red algae are effective indicators of detergent damage since they undergo colour changes when exposed to relatively low concentration of detergent. Smith (1968) reported that red algae such as Chondrus crispus, Mastocarpus stellatus, and Laurencia pinnatifida were amongst the algae least affected by detergents, whereas other species, including Lomentaria articulata were either killed or unhealthy. Smith (1968) reported that 10 ppm of the detergent BP 1002 killed the majority of specimens in 24hrs in toxicity tests. Laboratory studies of the effects of oil and dispersants on several red algal species concluded that they were all sensitive to oil/dispersant mixtures, with little difference between adults, sporelings, diploid or haploid life stages (Grandy, 1984; cited in Holt et al., 1995). Cole et al. (1999) suggested that herbicides were, not surprisingly, very toxic to algae and macrophytes. Hoare & Hiscock (1974) noted that all red algae except Phyllophora sp. were excluded from near to an acidified halogenated effluent discharge in Amlwch Bay, Anglesey and that intertidal populations of Corallina officinalis occurred in significant amounts only 600 m east of the effluent.

Most pesticides and herbicides were suggested to be very toxic for invertebrates, especially crustaceans (amphipods isopods, mysids, shrimp and crabs) and fish (Cole et al., 1999). For example, Lindane was shown to be very toxic to gobies (Gobius spp.: see Pomatoschistus minutus) (Ebere & Akintonwa, 1992). The pesticide ivermectin is very toxic to crustaceans, and has been found to be toxic towards some benthic infauna such as Arenicola marina (Cole et al., 1999). The evidence suggests that, on balance, red algae are probably very intolerant to synthetic chemicals and biotope intolerance is assessed as high. Recoverability is probably also high (see additional information below).

Heavy metal contamination
Low Immediate Not sensitive Minor decline Low
Bryan (1984) suggested that the general order for heavy metal toxicity in seaweeds is: organic Hg > inorganic Hg > Cu > Ag > Zn > Cd >Pb. Cole et al. (1999) reported that Hg was very toxic to macrophytes. The sub-lethal effects of Hg (organic and inorganic) on the sporelings of an intertidal red algae, Plumaria elegans, were reported by Boney (1971). 100% growth inhibition was caused by 1 ppm Hg. Burdin & Bird (1994) reported that both gametophyte and tetrasporophyte forms of Chondrus crispus accumulated Cu, Cd, Ni, Zn, Mn and Pb when immersed in 0.5 mg/l solutions for 24 hours. No effects were reported however, and no relationship was detected between hydrocolloid characteristics and heavy metal accumulation.

Cole et al. (1999) suggested that Pb, Zn, Ni and As were very toxic to algae, while Cd was very toxic to Crustacea (amphipods, isopods, shrimp, mysids and crabs), and Hg, Cd, Pb, Cr, Zn, Cu, Ni, and As were very toxic to fish. Bryan (1984) reported sublethal effects of heavy metals in crustaceans at low (ppb) levels. In laboratory investigations Hong & Reish (1987) observed 96hr LC50 (the concentration which produces 50% mortality) of between 0.19 and 1.83 mg/l in the water column for several species of amphipod. The intolerant of crustaceans to heavy metal contaminants suggests that amphipod and isopod grazers would be lost, allowing rapid growth of epiphytes, and perhaps reduced growth of Corallina officinalis.

In the absence of evidence of mortalities in red or coralline algae and intolerance of low has been recorded to represent the potential loss of epiphytic grazers, albeit with very low confidence.
Hydrocarbon contamination
High High Moderate Major decline Moderate
Oil and detergent dispersants affected high water specimens of Corallina officinalis more than low shore specimens and some specimens were protected in deep pools. In areas of heavy spraying, however, Corallina officinalis was killed, and was affect down to 6 m in one site, presumably due to wave action and mixing (Smith, 1968). However, regrowth of fronds had begun within 2 months after spraying ceased (Smith, 1968). Crump et al. (1999) noted a dramatic bleaching of encrusting corallines and signs of bleaching in Corallina officinalis, Chondrus crispus and Mastocarpus stellatus at West Angle Bay, Pembrokeshire after the Sea Empress oil spill. However, encrusting corallines recovered quickly and Corallina officinalis was not killed. It seems likely, therefore, that Corallina officinalis was more intolerant of dispersants used during the Torrey Canyon oil spill than the oil itself.

O'Brien & Dixon (1976) suggested that red algae were the most intolerant group of algae to oil or dispersant contamination, possibly due to the susceptibility of phycoerythrins to destruction. Laboratory studies of the effects of oil and dispersants on several red algal species concluded that they were all sensitive to oil/dispersant mixtures, with little difference between adults, sporelings, diploid or haploid life stages (Grandy, 1984; cited in Holt et al., 1995). Smith (1968) reported that red algae such as Chondrus crispus, Mastocarpus stellatus, and Laurencia pinnatifida were amongst the algae least affected by detergents, whereas other species, including Lomentaria articulata were either killed or unhealthy.

Suchanek (1993) noted that gastropods, amphipods, infaunal polychaetes and bivalves were particularly sensitive to oil spills. Amphipods in particular are known to be sensitive to oil spills (Suchanek, 1993).

Overall, hydrocarbon contamination and oil spills are likely to reduce the cover of the dominant macrophytes in the biotope, due to bleaching and subsequent loss of fronds. But the resistant bases of Corallina officinalis will probably survive and facilitate recovery. Some of the dominant fleshy red algae are likely to survive while other may be killed. But the epiphytic fauna is likely to be adversely affect due to loss of habitat, smothering by oil, and direct effects of oil contamination on amphipods, and other crustaceans, limpet and gastropods in particular. Therefore, an intolerance of intermediate has been recorded, while species richness is likely to decline markedly. Once the oil has been removed, which is probably rapid in a wave exposed habitat, subsequent recovery is probably rapid.

Radionuclide contamination
No information No information No information Insufficient
information
Not relevant
No information found.
Changes in nutrient levels
Low Immediate Not sensitive Minor decline Low
Corallines seem to be tolerant and successful in polluted waters. Kindig & Littler (1980) demonstrated that Corallina officinalis var. chilensis in South California showed equivalent or enhanced health indices, highest productivity and lowest mortalities (amongst the species examined) when exposed to primary or secondary sewage effluent. Little difference in productivity was noted in chlorinated secondary effluent or pine oil disinfectant. However, specimens from unpolluted areas were less tolerant, suggesting physiological adaptation to sewage pollution (Kindig & Littler, 1980).

Johansson et al. (1998) suggested that one of the symptoms of large scale eutrophication is the deterioration of benthic algal vegetation in areas not directly affected by land-runoff or a point source of nutrient discharge. Altered depth distributions of algal species caused by decreased light penetration (turbidity) and/or increased sedimentation through higher pelagic production have been reported in the Baltic Sea (Kautsky et al., 1986; Vogt & Schramm, 1991). An increase in abundance of red algae, including Delesseria sanguinea, was associated with eutrophication in the Skagerrak area, Sweden, especially in areas with the most wave exposure or water exchange (Johansson et al., 1998). However, where eutrophication resulted in high siltation rates, the delicate foliose red algae such as Delesseria sanguinea were replaced by tougher, erect red algae (Johansson et al., 1998).

Eutrophication in the intertidal is likely to favour ephemeral algae such as Ulva spp., Ulva lactuca and Porphyra spp., which in turn may favour epiphytic grazers. However, The coralline turf will probably survive and may even benefit from nutrient enrichment, and although the dominant red algae may change, favouring fast growing species, the biotope will probably remain recognizable. Therefore, an intolerance of low has been recorded.
Tolerant Not relevant Not relevant Insufficient
information
Low
This biotope occurs in the intertidal of fully saline waters, and organisms will be exposed to increased salinities due to evaporation during emersion. In addition, Corallina officinalis inhabits rock pools and gullies from mid to low water. Therefore, it is likely to be exposed to short term hypersaline (evaporation) events. Kinne (1971) cites maximal growth rates for Corallina officinalis between 33 and 38 psu in Texan lagoons. Overall, little information on hypersaline tolerance of other species was found, however it appear that the coralline turf will probably survive, and the biotope has been assessed as not sensitive, albeit at low confidence.
Intermediate Very high Low Decline Low
This biotope occur in the intertidal of fully saline waters. Corallina officinalis is likely to be exposed to short term hyposaline (freshwater runoff and rainfall) and hypersaline (evaporation) events where it occurs in rock pools. But the distribution of Corallina officinalis in the Baltic is restricted to increasingly deep water as the surface salinity decreases, suggesting that it requires full salinity in the long term (Kinne, 1971), although the coralline turf communities described by Bamber 1993) occurred in waters between 24 and 28psu. Gessner & Schramm (1971) summarize the effects of salinity changes on marine algae. Most sublittoral red algae cannot withstand salinities below 15 psu. Therefore, a reduction in salinity in the long term, from full to reduced is likely to result in a decline in the abundance of the coralline turf and red algae below about 24 psu, and hence its associated community. Therefore, an intolerance of intermediate has been recorded. Recoverability is likely to be very high.
Not relevant Not relevant Not relevant Not relevant Not relevant
This biotope occurs in wave exposed conditions with considerable mixing of the water column and wave crash. Therefore, hypoxic or anoxic conditions are unlikely to occur.

Biological Pressures

 IntoleranceRecoverabilitySensitivityRichnessEvidence/Confidence
No information Not relevant No information Insufficient
information
Not relevant
Several coralline and non-coralline species are epiphytic on Corallina officinalis. Irvine & Chamberlain (1994) cite tissue destruction caused by Titanoderma corallinae. However, no information on pathogenic organisms in the UK was found. In Rhodophyta, viruses have been identified by means of electron microscopy (Lee, 1971) and it is obvious that they are widespread. But nothing is known of their effects on growth or reproduction in red algae and experimental transfer from an infected to an uninfected specimen has not been achieved (Dixon & Irvine, 1977). Overall, insufficient information was found to make an assessment.
No information Not relevant No information Insufficient
information
Not relevant
No information found.
Intermediate Very high Low Decline Low
Corallina officinalis was used in Europe as a vermifuge although it no longer seems to be collected for this purpose (Guiry & Blunden, 1991). Corallina officinalis is collected for medical purposes; the fronds are dried and converted to hydroxyapatite and used as bone forming material (Ewers et al., 1987). It is also sold as a powder for use in the cosmetic industry. An European research proposal for cultivation of Corallina officinalis is pending as of May 2000 (Wiedemann, pers. comm.). Both Chondrus crispus and Mastocarpus stellatus are collected as 'carragheen' by hand picking and racking in Europe (Guiry & Blunden, 1991). Removal of any of the macroalgal community would obviously reduce its extent and cover but also significantly reduce the resident epiphytic fauna. Intolerance has been assessed as intermediate. However, as long as holdfasts remain recovery will probably be rapid.
Not relevant Not relevant Not relevant Not relevant Not relevant

Additional information

Recoverability
Corallina officinalis probably has good recruitment and settled on artificial substrata within 1 week of their placement in the intertidal during summer in New England (Harlin & Lindbergh, 1977). New fronds of Corallina officinalis appeared on sterilised plots within six months and 10% cover was reached with 12 months (Littler & Kauker, 1984). Bamber (1993) reported that new plants grew back in scraped transects within 12 months, although the resistant crustose bases were probably not removed. Similarly, in experimental plots, up to 15% cover of Corallina officinalis fronds returned within 3 months after removal of fronds and all other epiflora/fauna but not the crustose bases (Littler & Kauker, 1984). Although new crustose bases may recruit and develop quickly the formation of new fronds from these bases and recovery of original cover may take longer, and it is suggested that a the population is likely to recover within a few years.

If the holdfasts of red algae remain, they are likely to recover quickly, as if damaged by winter storms. For example, following experimental harvesting by drag raking in New Hampshire, USA, populations of Chondrus crispus recovered to one third of their original biomass after 6 months and totally recovered after 12 months (Mathieson & Burns, 1975). Raking is designed to remove the large fronds but leave the small upright shoots and holdfasts. The authors suggested that control levels of biomass and reproductive capacity are probably re-established after 18 months of regrowth, although time to recovery was much extended if harvesting occurred in the winter, rather than the spring or summer (Mathieson & Burns, 1975). Minchinton et al. (1997) documented the recovery of Chondrus crispus after a rocky shore in Nova Scotia, Canada, was totally denuded by an ice scouring event. Initial recolonization was dominated by diatoms and ephemeral macroalgae, followed by fucoids and then perennial red seaweeds. After 2 years, Chondrus crispus had re-established approximately 50% cover on the lower shore and after 5 years it was the dominant macroalgae at this height, with approximately 100% cover. Therefore, recovery by Chondrus crispus will be relatively rapid (approximately 18 months) in situations where intolerance to a factor is intermediate and some holdfasts remain for regeneration of fronds. In situations of high intolerance, where the entire population of Chondrus crispus is removed, recovery will be limited by recruitment from a remote population and would be likely to take up to 5 years. Similarly, clearance studies of concrete blocks in the shallow subtidal showed that Rhodophyceae colonized and grew in the winter months, presumably at their peak of spore availability (Kain, 1975). It is probably that most of the characterizing red algae would grow back from remaining holdfasts and recruit well in winter and where bases are removed they will probably take a few years to regain their original cover, although in this biotope their percentage cover is low.

The epiphytic fauna are mainly composed of mobile species, that will recruit quickly from surrounding habitats, and will therefore, recover quickly once the coralline turf has developed.

Overall, where upright fronds of the red algal turf are removed, recovery will probably be very rapid, within about 12 months. If the holdfasts are removed, recovery of their original cover is likely to be prolonged but the biotope would probably be recognizable within less than 5 years.

Importance review

Policy/Legislation

Habitats Directive Annex 1Reefs

Exploitation

Corallina officinalis was used in Europe as a vermifuge although it no longer seems to be collected for this purpose (Guiry & Blunden, 1991). Corallina officinalis is collected for medical purposes; the fronds are dried and converted to hydroxyapatite and used as bone forming material (Ewers et al., 1987). It is also sold as a powder for use in the cosmetic industry. An European research proposal for cultivation of Corallina officinalis is pending as of May 2000 (Wiedemann, pers. comm.). Both Chondrus crispus and Mastocarpus stellatus are collected as 'carragheen' by hand picking and racking in Europe (Guiry & Blunden, 1991).

Additional information

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Citation

This review can be cited as:

Tyler-Walters, H., 2005. Corallina officinalis on exposed to moderately exposed lower eulittoral rock. In Tyler-Walters H. and Hiscock K. (eds) Marine Life Information Network: Biology and Sensitivity Key Information Reviews, [on-line]. Plymouth: Marine Biological Association of the United Kingdom. Available from: http://www.marlin.ac.uk/habitat/detail/130

Last Updated: 12/01/2005